Revision of the poorly known Neotropical butterfly genus Zischkaia Forster, 1964 (Lepidoptera, Nymphalidae, Satyrinae), with descriptions of nine new species Author Nakahara, Shinichi A7240997-442C-41C7-9B20-D8D4AE31AED9 McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA. & Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611, USA. & Departamento de Entomología, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru. snakahara@ufl.edu Author Zacca, Thamara EBE99273-B02D-4A17-AD52-5BB3DFBA6C85 Departamento de Biologia Animal and Museu de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, Campinas, São Paulo, Brazil. zacca.butterfly@gmail.com Author Dias, Fernando M. S. FA0950D9-DC58-4230-8180-F26CB5788E55 Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. fernandomsdias@yahoo.com.br Author Dolibaina, Diego R. 0A762FB6-AF41-4A09-9DF8-6F3954BA8956 Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. dirodrido@gmail.com Author Xiao, Lei 4233F498-4D30-428A-B2A4-A6A76DC5FCA4 McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA. lxiao@floridamuseum.ufl.edu Author Espeland, Marianne 00D6F9F9-3902-4A8B-846F-720AB32922A6 Arthropoda Department, Zoological Research Museum Alexander Koenig, Adenauer Allee 160, 53113 Bonn, Germany. marianne.espeland@gmail.com Author Casagrande, Mirna M. Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. mibras@ufpr.br Author Mielke, Olaf H. H. Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. omhesp@ufpr.br Author Lamas, Gerardo Departamento de Entomología, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru. glamasm@unmsm.edu.pe Author Huertas, Blanca Life Sciences Department, Natural History Museum, London, UK. B.Huertas@nhm.ac.uk Author Kleckner, Kaylin McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA. & Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611, USA. kaylin.kleckner@ufl.edu Author Willmott, Keith R. McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA. kwillmott@flmnh.ufl.edu text European Journal of Taxonomy 2019 2019-09-13 551 1 67 journal article 10.5852/ejt.2019.551 98f587bb-3baf-4961-89a0-5679c0dcf731 2118-9773 3476676 C3C851C3-0F12-412C-A15B-56F0F263CD00 Zischkaia pacarus (Godart, [1824]) Figs 1 , 2 S–V, 4 G, 6 G–H, 8 A, 10 A–C, 11 , 12 A–D, 14 Satyrus pacarus Godart, [1824]: 465, 495 ( type locality: Brazil ). Euptychia fumata Butler, 1867b: 109 , pl. 12, fig. 14 ( type locality: “Rio Grande”). Neonympha pacarus – Westwood 1851: 375. Euptychia pacarus Butler 1867a: 501 ; 1868: 38 ; 1877: 120 . — Kirby 1871: 55 . — Gaede 1931: 458 . — Weymer 1911: 213 . — D’Abrera 1988: 789 . Euptychia fumata Butler 1867a: 501 , nomen nudum. Euptychia fumata Butler 1868: 38 ; 1877: 120 . — Kirby 1871: 55 ; 1879: 134 . — Weymer 1911: 213 . — Riley & Gabriel 1924: 22 . — Gaede 1931: 446 . — Miller 1968: 95 . — D’Abrera 1988: 789 . Zischkaia fumata Forster 1964: 116–118 , fig. 123 (the description date of the name was mistakenly given as “1857”). — Brown 1992: 152 , fig. 49. — Mielke & Casagrande 1998: 976 . — Brown & Freitas 2000: 105 . Zischkaia pacarus Mielke 1995: 772 . — Lamas 2004: 223 , syn.: fumata Butler, 1867 . — Emery et al. 2006: 90 . — Beccaloni et al. 2008: 346 . — Carneiro et al. 2008: 264 . — Peña et al. 2010: 248 , 250–251, 253. — Dolibaina et al. 2011: 350 . — Silva et al. 2015: 6 . Type material examined Neotype (here designated) BRAZIL ; “ // BRASIL, PARANÁ , TUNEIRAS DO OESTE , REBIO [ Reserva Biológica ] DAS PEROBAS , 23° 50” 49”S 52° 44” 18” W , 8-X-2012 LABLEP [ Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná] LEG.// BC-DZ Willmott 246// DZ 36.998//”; DZUP . Lectotype (here designated) BRAZIL ; “(// Rio Grande [do Sul, Brazil] Hewitson coll[ection]. 79-69 Euptychia Fumata, Butl [er]. 2.// BM TYPE No. Rh 3251 Euptychia fumata Butl[er].// Type // BMNH(E) 1267093//”; NHMUK . Other material examined ( 87 ♂♂ , 50 ♀♀ ) ARGENTINA Misiones 2 ♂♂ ; Apr. 1927 ; I. Heider leg.; BMNH(E)-10247985, BMNH(E)-10430700; NHMUK 1 ♀ ; El Dorado ; [ 26°23′ S , 54°40′ W ]; ZSM 1 ♂ ; Loreto ; [ 27°19′ S , 55°32′ W ]; Oct. 1956 ; W. Forster leg.; ZSM . BOLIVIA Not located 1 ♂ ; “Torochila” [perhaps representing “Jorochito”]; Garlepp leg.; 1890; MNHU . BRAZIL Bahia 1 ♂ ; Una , Reserva Biológica de Una ; [ 15°10′38″ S , 39°6′20″ W ]; G.M. Accácio leg.; ZUEC . – Espírito Santo 1 ♂ ; Fruhstorfer leg.; BMNH(E)-1205412; NHMUK . – Minas Gerais 1 ♀ ; S of Minas Gerais ; BMNH(E)-1718052; NHMUK 2 ♀♀ ; Carangola , Serra Rio Glória , Sítio Boa Vista ; [ 20°43′44″ S , 42°3′27″ W ]; 600–800 m ; Dec. 1950 ; Ebert leg.; DZ-36529, DZ-36549; DZUP 4 ♂♂ ; Porto Real ; [ 20°11′ S , 45°43′ W ]; BMNH(E)-1718043 to BMNH(E)-1718045, BMNH(E)-1718056; NHMUK 2 ♀♀ ; same collection data as for preceding; BMNH(E)-1718050 to BMNH(E)-1718051; NHMUK 1 ♂ ; San [São] Jacinto ; [ 17°51′ S , 41°30′ W ]; Nov. 1907 ; F. Birch leg.; BMNH(E)-1718042; NHMUK . – Paraná 1 ♂ ; BMNH(E)-1205411]; NHMUK 1 ♀ ; BMNH(E)-1718062; NHMUK 1 ♂ ; Balsa Nova , São Luís do Purunã ; 1000 m ; 8 Apr. 2006 ; Beltrami and Selusniaki leg.; DZ-36643; DZUP 1 ♂ ; Campo Mourão , ( Justus ); Jan. 1952 ; DZ-36861; DZUP 5 ♂♂ ; Candói , Santa Clara ; [ 25°37′ S , 51°59′ W ]; 21 Nov. 1986 ; O.H.H. Mielke and M.M. Casagrande leg.; DZ-36589, DZ-36633, DZ-36831, DZ-36891, DZ-5575; DZUP 2 ♀♀ ; same collection data as for preceding; DZ-36851, DZ-36991; DZUP 1 ♂ , 1 ♀ ; Castro ; [ 24°47′ S , 50°1′ W ]; 950 m ; E.D. Jones leg.; BMNH(E)-1718058, BMNH(E)-1718055; NHMUK 1 ♂ ; same collection data as for preceding but collector unknown; BMNH(E)-1718041; NHMUK 2 ♂♂ ; Chopinzinho ; [ 25°51′ S , 52°30′ W ]; 1 Mar. 1970 ; E. Furtado leg.; DZ-36971, DZ-37031; DZUP 1 ♂ ; Foz do Iguaçu ; [ 25°30′ S , 53°48′ W ]; Mar. 1952 ; Justus leg.; DZ-37021; DZUP 1 ♂ ; Iguazu ; 24 Jan. 1922 ; BMNH(E)-1718057; NHMUK 1 ♀ ; Londrina ; [ 23°19′ S , 51°10′ W ]; 25 Oct. 1985 ; O.H.H. Mielke and M.M. Casagrande leg.; DZ-37041; DZUP 1 ♀ ; Ponta Grossa ; [ 25°6′ S , 50°10′ W ]; Nov. 1956 ; Justus leg.; DZ-36821; DZUP 1 ♀ ; Vila Velha ; [ 25°8′ S , 49°58′ W ]; 900 m ; 16 Mar. 1972 ; Mielke leg.; DZ-36841; DZUP 1 ♀ ; 7 Mar. 1973 ; DZ-36981; DZUP 1 ♀ ; Lapa , Rio de Várzea ; [ 25°46′ S , 49°43′ W ]; Feb. 1938 ; Pohl leg.; MZSP 1 ♀ ; Tibagi , Parque Estadual do Guartelá , [ 24°33′59″ S , 50°15′25″ W ]; 1000 m ; 11–12 Feb. 2016 ; D. Dolibaina , E. Carneiro and A. Warren leg.; DD-374 6 ♂♂ , 4 ♀♀ ; Tuneiras do Oeste , Reserva Biológica das Perobas ; [ 23°50′49″ S , 52°44′18″ W ]; (‘LELN UFPR’); 6 Oct. 2012 ; DZ-36498, DZ-36728, DZ-36858, DZ-36918, DZ-36939, DZ-36949, DZ-36778, DZ-36909, DZ-36919, DZ-36959; DZUP 4 ♂♂ , 2 ♀♀ ; same collection data as for preceding but 8 Oct. 2012 ; DZ-36708, DZ-36929, DZ-36989, DZ-37039, DZ-36588, DZ-36999; DZUP 1 ♂ ; [ 23°53′ S , 52°44′ W ; 578 m ; 6 Oct. 2012 ; G. Lamas leg.; MUSM 1 ♂ ; same collection data as for preceding but 8 Oct. 2012 ; MUSM 1 ♂ ; Turvo , Britador ; [ 25°2′11″ S , 51°32′8″ W ]; 1000 m ; 19 Feb. 2010 ; D. Dolibaina and A. Warren leg.; DD-303 1 ♂ , 1 ♀ ; Turvo , Salto do Paulinho Rickli ; [ 25°3′59″ S , 51°31′8″ W ]; 1050 m ; 15 Mar. 2009 ; D. Dolibaina leg.; DD-301 to DD-302 4 ♂♂ , 1 ♀ ; same collection data as for preceding but 7 Feb. 2016 ; D. Dolibaina and Warren leg.; DD-369 to DD-373 . – Rio Grande do Sul 1 ♀ ; 1890; Mabilde leg.; MNHU 1 ♀ ; Elsenau [Panambi]; Dec. 1904 ; ZSM 3 ♂♂ , 2 ♀♀ ; Campo Novo ; 9 Nov. 1985 ; Araújo, O.H.H . Mielke and M.M. Casagrande leg.; DZ-36781, DZ-36941, DZ-36951, DZ-36741, DZ-36751; DZUP 2 ♀♀ ; Derrubadas , Parque Florestal Estadual do Turvo ; [ 27°13′58″ S , 53°51′5″ W ]; 10 Nov. 1985 ; O.H.H. Mielke , Araújo , and M.M. Casagrande leg.; DZ-36761, DZ-36811; DZUP 1 ♀ ; Pelotas ; [ 31°45′ S , 52°22′ W ]; 15 Oct. 1953 ; C.M. Biezanko leg.; DZ-36931; DZUP 1 ♂ ; 28 Feb. 1961 ; AMNH 1 ♂ , 2 ♀♀ ; 11 Mar. 1961 ; USNM 1 ♂ ; 25 Nov. 1961 ; AMNH 1 ♂ ; 12 Dec. 1961 ; USNM 1 ♀ ; 22 Mar. 1964 ; AMNH 1 ♂ ; 13 Feb. 1966 ; V. Becker leg.; DZ-36801; DZUP 1 ♀ ; 6 Nov. 1961 ; C.M. Biezanko leg.; ZSM 1 ♂ ; 7 Nov. 1966 ; ZSM 1 ♂ ; 7 Nov. 1966 ; DZ-36971; DZUP 1 ♂ ; 3 Mar. 1967 ; DZ-36791; DZUP 1 ♂ ; 7 Mar. 1970 ; DZ-21207; DZUP . – Santa Catarina 1 ♂ ; Wernicke leg.; ZSM 1 ♂ ; Rio Capivary [Capivari]; 1883; Fruhstorfer leg.; BMNH(E)-1718059; NHMUK 1 ♀ ; ridge between Joinville and Itajaí , Tajú ; [ 26°34′26″ S , 48°44′51″ W ]; Apr. ; Schmid leg.; DZ-36881; DZUP 1 ♂ ; same collection data as for preceding but Feb. ; DZ-37001; DZUP 1 ♂ ; Águas Mornas , Teresópolis ; [ 27°44′ S , 48°56′ W ]; BMNH(E)-1718048; NHMUK 1 ♀ ; same collection data as for preceding; Michalis leg.; BMNH(E)-1718063; NHMUK 1 ♂ ; Blumenau ; [ 26°55′ S , 49°4′ W ]; Fruhstorfer leg.; BMNH(E)-1205413; NHMUK 1 ♀ ; Florianópolis , Naufragados ; 11 May 2003 ; E. Carneiro leg.; DZ-37011; DZUP 5 ♂♂ , 1 ♀ ; Seara , Nova Teutonia ; [ 27°3′ S , 52°23′ W ]; 300–500 m ; Feb. 1973 ; F. Plaumann leg.; FLMNH-MGCL-1036000 to FLMNH-MGCL-1036005; FLMNH 1 ♂ ; 1 Feb. 1972 ; F. Plaumann and O.H.H. Mielke leg.; DZ-36658; DZUP 1 ♂ ; Taió , Rio Taió ; [ 27°6′ S , 49°59′ W ]; 2 Oct. 1935 ; Pohl leg.; MZSP 1 ♀ ; Timbó ; [ 26°49′ S , 49°17′ W ]; Apr. 1936 ; Pohl leg.; MZSP . – São Paulo 1 ♀ ; Araras ; [ 22°19′ S , 47°58′ W ]; 600 m ; 17 Apr. 1966 ; Ebert leg.; DZ-36519; DZUP 1 ♂ , 1 ♀ ; same collection data as for preceding but 6 Jan. 1966 ; DZ-36499, DZ-5577; DZUP 1 ♂ ; Casa Branca ; [ 21°47′ S , 47°5′ W ]; G[arbe] leg.; MNHU 1 ♂ [conf]; 1893–1895 ; Garbe leg.; MNHU 1 ♂ ; Cotia , Morro Grande ; 22 Nov. 2000 ; K.S. Brown and A.V.L. Freitas leg.; ZUEC 1 ♂ ; Guararapes , Figueira ; 5 Apr. 1940 ; R.F. d’Almeida leg.; DZ-36509; DZUP 1 ♀ ; Paranapiacaba , Alto da Serra ; Feb. 1922 ; Spitz leg.; MZUSP-56158; MZSP 1 ♂ ; Rio Claro , [ 22°24′ S , 47°33′ W ]; 600 m ; 13 Apr. 1967 ; Ebert leg.; DZ-36911; DZUP 1 ♀ ; same collection data as for preceding but 26 May 1963 ; DZ-5576; DZUP 1 ♂ , 1 ♀ ; same collection data as for preceding but 31 Dec. 1966 ; DZ-5578, DZ-36559; DZUP 1 ♂ ; Rio das Cobras ; [ 23°7′13″ S , 46°5′44″ W ]; Feb. 1942 ; Widerski leg.; DZ-36539; DZUP 1 ♀ ; Salesópolis , Estação Biológica da Boraceía ; [ 23°40′ S , 45°53′ W ]; 850 m ; 20 Mar. 1968 ; Lopes leg.; MZSP 1 ♂ ; same collection data as for preceding but 27 Feb. 1968 ; Oliveira Santos leg.; MZSP 1 ♂ ; São Paulo ; [ 23°32′ S , 46°37′ W ]; 790 m ; BMNH(E)-1718047; NHMUK 3 ♂♂ , 1 ♀ ; same collection data as for preceding; Garbe leg.; MZUSP-56159 to MZUSP-56162; MZSP 1 ♂ , 1 ♀ ; Teodoro Sampaio , Parque Estadual Morro do Diabo ; [ 22°30′ S , 52°20′ W ]; 250–500 m ; 15–18 Mar. 1990 ; O.H.H. Mielke and M.M. Casagrande leg.; DZ-36569, DZ-36579; DZUP 1 ♂ ; same collection data as for preceding but 15–18 Mar. 1991 ; DZ-36901; DZUP 1 ♂ , 1 ♀ ; same collection data as for preceding but 20–23 Feb. 1990 ; DZ-36921, DZ-36613; DZUP 1 ♂ ; same collection data as for preceding but 22–23 May 1990 ; DZ-36771; DZUP . – No specific locality 1 ♂ ; H. Rogers leg.; PLT fumata ; BMNH(E)-1718068; NHMUK Not located 1 ♂ ; ‘ Brazil’ ; BMNH(E)-1718046; NHMUK 1 ♂ ; ‘ Brazil’ ; PLT fumata ; BMNH(E)-1718067; NHMUK 1 ♀ ; ‘ Brazil’ ; PLT fumata ; BMNH(E)-1718066; NHMUK . Fig. 10. Head morphology. A–C . Zischkaia pacarus ( Godart, 1824 ) (DZ 36.728). A . Ventral view. B . Frontal view. C . Lateral view. D–F . Z. saundersii ( Butler, 1867 ) (DZ 36.029). D . Ventral view. E . Frontal view. C . Lateral view. PARAGUAY Caaguazú 1 ♂ ; Caaguazú ; [ 25°26′ S , 56°2′ W ]; Apr. 1960 ; ZSM . VENEZUELA Not located 1 ♂ ; ‘Venezuela’, BMNH(E)-1718065; NHMUK . COUNTRY UNKNOWN – Not located1 ♂ ; no data; BMNH(E)-1718049; NHMUK . Identification and taxonomy The male of Zischkaia pacarus is distinguishable from males of other species of Zischkaia by the apical process of the valva being somewhat rectangular with a convex distal margin ( Fig. 4G ), whereas the apical process of the valva is somewhat triangular in other species of Zischkaia . The female of Z. pacarus can also be distinguished by the genitalia ( Fig. 6 G–H), namely by the sclerotized tube being one-fourth the length of the ductus bursae, whereas this sclerotized tube is reduced and restricted to the area around the ostium bursae in Z. chullachaki sp. nov. , more developed and reaching almost half the length of the ductus bursae in Z. baku sp. nov. and Z. arenisca sp. nov. Godart ([1824]: 495) described Satyrus pacarus based on a single specimen from Brazil , without any mention of the sex of this specimen. He provided a short description of the species (Godart ([1824]: 465), stating “[margin of the] wings entire, dark brown, upper side without spots; forewing under side with two darker wavy lines, hind wing under side with three [darker wavy lines], and five eyes [i.e., ocelli] having the pupil formed by silver dots.” The full description on page 495 is lengthier and more precise: “It has approximately two inches [about 5 cm ] of wingspan. Its wings [margins] are entire, and dark brown in both sides. Its [wings] upper sides are without spots. Its [wings] under sides with, towards the extremity, two lines darker than the ground color. The second wings [i.e., hindwings] have a third similar line, placed transversely towards the middle, and separated from the others by a row of five round, very dark, eyes [i.e., ocelli], with a pupil formed by silver dots, and a brownish iris surrounded by a very dark circle. The first wings [forewings] have no eyes [i.e., ocelli].” (translation based on the original description of both French and Latin version). Certain phenotypic characters described by Godart, namely, no mention of the VFW ocellus (or ocelli), no ocellus (or ocelli) on the dorsal surface, the mention of two lines on the VFW and three lines on the VHW (i.e., lack of VFW submedian line), and the fact this species was allied with several other euptychiine species (e.g., Hermeuptychia sosybius (Fabricius, 1793)) , suggest that Godart was describing a species of Zischkaia . There is no mention of dorsal surface androconia in Godart’s description, one of the putative synapomorphic characters for the “ pacarus clade” of Zischkaia , and we offer two possible explanations: 1) although the specimen he examined was a male, Godart failed to perceive or did not regard as important the presence of this character; 2) the specimen examined by Godart was a female, and therefore without androconia. Given that the dorsal androconia are not well marked in Z. pacarus , and considering that females are rarer than males, the former is perhaps the more likely. For example, Godart also did not describe the androconial patch of Calisto hysius hysius (Godart, [1824]) (described on page 471) and Forsterinaria necys (Godart, [1824]) (described on page 511), although males of these two species also possess dorsal wing androconial patches. There remains a possibility that Godart was describing Z. warreni sp. nov. , which also lacks a VFW submedian line and occurs in sympatry with Z. pacarus . However, Z. warreni sp. nov. has a much narrower geographic range and is much scarcer than Z. pacarus in museum collections ( 43 specimens examined of the former species and 138 of the latter). Most likely, the holotype (fixed by monotypy, following Article 73.1.2. of the ICZN (1999)) of pacarus should have been collected in Minas Gerais or Rio de Janeiro , where Z. warreni sp. nov. is apparently rare (see below). Furthermore, at least some of the species described by Godart are not known or are very rare within the range of Z. warreni sp. nov. (e.g., Godartiana byses (Godart, [1824]), see Zacca et al. (2017) , Adelpha epione (Godart, [1824]), see Willmott (2003)) , suggesting that the type of Z. pacarus might have came from further east in southern Brazil . We therefore contend that Godart’s description most likely referred to Z. pacarus as treated here. The holotype of S. pacarus has not been located in any museums where specimens from Godart’s collection are usually found (e.g., MNHN ; Royal Scottish Museum, Edinburgh ( Grimshaw 1897 ; Horn et al. 1990 ; Bland 2019 )). Therefore, a male specimen deposited at the DZUP ( Fig. 2 ), which we also barcoded and included in the phylogeny ( BC-DZ 246), is designated as the neotype of Satyrus pacarus Godart , [1824] in order to settle the nomenclature ( neotype designation). Fig. 11. Zischkaia pacarus ( Godart, 1824 ) , wing venation. A . ♂ (FLMHN-MGCL-1036002). B . ♀ (MGCL-LOAN 518). Fig. 12. Wing scales of Zischkaia Forster, 1964 . A–C . Images of forewing androconial scales of Z. pacarus (Godart, [1824]), showing the presence of plumose androconial scales. D . Typical wing scales of Z. pacarus . E–F . Images of forewing androconial scales of Z. chullachaki Nakahara & Zacca , sp. nov. , showing the presence of plumose androconial scales. Butler (1867a) introduced the name Euptychia fumata , but in a way that failed to fulfill Article 12.1 of the ICZN (1999) , as the name was not accompanied by a description, definition, or indication of the taxon it denotes. Therefore, the name was correctly recognized as a nomen nudum by Lamas (2004) . Subsequently, Butler (1867b) properly described Euptychia fumata Butler, 1867 based on an unknown number of specimens from “Rio Grande” (likely the state of Rio Grande do Sul , southern Brazil ) from Hewitson’s collection. Given the presence of other similar Zischkaia species in southern Brazil , a male syntype , deposited in the NHMUK , which compares closely to the description and illustration provided by Butler (1867b) , is here designated lectotype , in order to settle the nomenclature ( lectotype designation). Euptychia fumata Butler, 1867 was treated as a junior subjective synonym of Z. pacarus by Lamas (2004) . Although Lamas (2004) did not provide any justification for this synonymy, the lectotype of E. fumata , here designated, compares closely with Godart’s description of S. pacarus and the neotype male designated above; therefore, we retain Euptychia fumata Butler, 1867 as a junior subjective synonym of Z. pacarus . Variation The conspicuous dorsal androconia is variable, from being slightly to moderately marked; the VHW ocellus in Rs-M 1 is variable in terms of presence and development; the visibility of the bright silvery purple ground color between the VHW submedian line and submarginal line when seen under direct light is also apparently variable, from absent to present. Distribution ( Fig. 14 ) This species is known from southwestern and southern Brazil to northeastern Argentina , in the states of Minas Gerais , São Paulo , Paraná , Santa Catarina , Rio Grande do Sul ( Brazil ), in the department of Caaguazú ( Paraguay ) and the province of Misiones ( Argentina ). Old records from Amazon Superior, Brazil , Bolívia and Venezuela probably refer to mislabeled specimens. Remarks In Araucaria forests in Paraná , adults of Z. pacarus are found during the summer either on small and isolated or large and continuous patches of the bamboo Merostachys Spreng. ( Poaceae : Bambusoideae ) ( Fig. 16A, C ). Males fly fast, up to 5 m above the ground, with several individuals engaged in patrolling flight. Females fly slowly and lower than males. Butterflies are never found away from the bamboo, and they are more active between the middle of the morning and the beginning of the afternoon. The preferred habitat of Z. pacarus in Araucaria forests suggests that the species likely uses Merostachys as the caterpillar host plant. In addition, according to A.V.L Freitas (pers. comm. in Beccaloni et al. 2008 ), larvae feed on Bambusa ( Poaceae : Bambusoideae ).